Association patterns of sailfin mollies (Poecilia latipinna): alternative hypotheses

Individuals may associate with each other due to a variety of selective forces, such as intra- and intersexual selection, and conspecific recognition. Previous studies have concluded that mate choice governs association behavior in polygynous species of fish. I examined whether mate choice underlies the preference for larger individuals by examining preference for association (time spent in proximity to a fish) not only between opposite-sex individuals but also between same-sex individuals of the live-bearing sailfin molly (Poecilia latipinna). Males and females from three size classes were tested with a large and a small object fish of the same and opposite sex. Females preferred to associate with larger over smaller males. Males also preferred to associate with larger over smaller females, as expected. The same female and male test fish also preferred to associate with larger over smaller fish of the same sex. Moreover, females demonstrated no significant difference in their strength of preference (large–small) when offered males or females. The same held true for males. When males and females were subsequently tested with one large male and one large female, females tended to prefer large males while males showed no significant preference for association based on sex. In another experiment, females were tested with a large female and a small male, and significantly preferred the former. These findings suggest that association patterns may have arisen under a variety of conditions, such as predation pressures, shoaling behavior, and associative preference behavior. The assumption that association behavior is a uniformly sufficient predictor of mate choice in fish needs to be re-examined for P. latipinna and other species. Received: 6 November 1998 / Received in revised form: 12 May 1999 / Accepted: 12 May 1999     

The effect of perceived female parasite load on post-copulatory male choice in a sex-role-reversed pipefish

The last several decades of research in behavioral ecology have resulted in a deeper appreciation of post-mating processes and sexual conflict in sexual selection. One of the most controversial aspects of sexual selection is cryptic mate choice. Here, we take advantage of male pregnancy in a sex-role-reversed pipefish (Syngnathus typhle) to quantify cryptic choice based on perceived parasite load and other sources of variance in female fitness. Studies have shown that S. typhle males preferentially mate with females with lower parasite loads and that a male’s perception of female parasite load can be altered by tattooing females. We manipulated the apparent parasite load of females in controlled mating experiments to test the hypothesis that post-copulatory sexual selection is dependent on a male’s perception of female parasite load in pipefish. Our results provided no evidence for cryptic male choice based on perceived female parasite load. However, we found evidence that eggs from larger females were more likely to result in viable offspring than eggs from smaller females and that the first female to mate with a male transferred more eggs per copulation on average. Overall, our results show that potential for post-copulatory sexual selection does exist in pipefish, but the male’s perception of female parasite load does not play a major role in this process.     

The roles of echolocation and olfaction in two Neotropical fruit-eating bats, Carollia perspicillata and C. castanea, feeding on Piper

We studied the echolocation and foraging behavior of two Neotropical frugivorous leaf-nosed bats (Carollia perspicillata, C. castanea: Phyllostomidae) in a flight cage. To test which cues Carollia uses to detect, identify, and localize ripe Piper fruit, their preferred natural food, we conducted experiments under semi-natural conditions with ripe, unripe, and artifical fruits. We first offered the bats ripe fruits and documented their foraging behavior using multiflash stereophotography combined with simultaneous sound recordings. Both species showed a similar, stereotyped foraging pattern. In searchflight, the bats circled through the flight cage in search of a branch with ripe fruit. After finding such a branch, the bats switched to approach behavior, consisting of multiple exploration flights and the final approach when the bats picked up the fruit at its tip and tore it off in flight. Our behavioral experiments revealed that odor plays an important role in enabling Carollia to find ripe fruit. While foraging, Carollia always echolocated and produced multiharmonic, frequency-modulated (FM) signals of broad bandwidth, high frequency, short duration, and low intensity. We discriminated an orientation phase (mostly a single pulse per wingbeat) and an approach phase (groups of two to six pulses per wing beat). We conclude from the bats' behavioral reaction to real and artificial fruit as well as from characteristic patterns in their echolocation behavior that during exploration flights, Carollia changes from primarily odor-oriented detection and initial localization of ripe fruit to a primarily echo-oriented final localization of the position of the fruit. Received: 27 March 1997 / Accepted after revision: 28 February 1998     

Shell choice in<Emphasis Type="Italic"> Pagurus longicarpus</Emphasis> hermit crabs: does predation threat influence shell selection behavior?

Pagurus longicarpus hermit crabs depend on empty gastropod shells for protection against predation. Hermit crabs avoid gastropod shells in which holes have been drilled by naticid gastropods, and hermit crabs forced to occupy drilled shells are more vulnerable to predation by green crabs, Carcinus maenas. In this study, we examined the effect of predator cues on P. longicarpus shell investigation behavior and shell choice. In paired laboratory shell choice trials, we examined hermit crab response to green crab chemical cues. We compared hermit crabs from two sites differing in the percentage of Littorina littorea shells with drill holes. The percentage of time hermit crabs spent occupying intact shells increased significantly in the presence of predator cues. The effect of predator cues on the amount of time hermit crabs spent investigating shells differed between individuals from the two sites. Predator effluent had a marginal effect on the proportion of hermit crabs initially choosing intact shells and within 15 min most hermit crabs in both treatments occupied intact shells due to shell switching. These results indicate that predation cues alter P. longicarpus shell choice behavior favoring intact shells, which provide greater protection. In summary, predation appears to be a key factor influencing hermit crab shell selection behavior.Communicated by T. Czeschlik     

Learning affects host discrimination behavior in a parasitoid wasp

Learning is generally predicted not to be important in host discrimination by parasitoids, because the stimuli involved are less variable than those used in habitat location. However, Anaphes victus (Hymenoptera: Mymaridae), an egg parasitoid of Listronotus oregonensis (Coleoptera: Curculionidae) apparently learns to associate external pheromones with the presence of a conspecific in a host. In this species, females can reject a parasitized host either after antennal drumming (antennal rejection) or after the insertion of their ovipositor (sting rejection). When they encountered a series of parasitized hosts, females A. victus learned to associate the presence of the external pheromone with the presence of the internal one. Learning lasted less than 4 h and occurred earlier in a series when the female marking the egg and the one detecting that mark were close relatives. This behavior could be adaptive because antennal rejection is faster than sting rejection. Received: 11 March 1997 / Accepted after revision: 30 August 1997     

Mating behavior of female rock shrimp Rhynchocinetes typus (Decapoda: Caridea)—indication for convenience polyandry and cryptic female choice

While studies on a wide diversity of organisms have demonstrated the importance of female behavior during matings, in crustacean studies, a strong bias towards male mating behavior prevails. Reproductively mature rock shrimp (Rhynchocinetes typus) exist as several ontogenetic stages that differ in their morphological and physiological capacities. In natural populations, the majority of males are in early ontogenetic stages (termed typus), many are in intermediate stages (intermedius), and few are in the terminal molt stage (robustus). Dominant robustus males, which have already demonstrated their biological fitness by surviving to this stage, have previously been shown to have a higher potential than subordinate typus males to defend receptive females against other males, and fertilize the entire clutch of a female. While females should thus show a preference for robustus males, they nevertheless frequently receive sperm from typus males. These observations suggested that females might have mechanisms to discriminate against sperm from subordinate males. In laboratory experiments, we observed that females avoided being seized by typus males for longer time periods in the absence of robustus males than in their presence. Following seizure, females that were initially held by typus males, required more time to initiate spawning than those held by robustus males. Many typus males transferred spermatophores to females before these started to spawn while robustus males waited until females began to spawn before they transferred spermatophores. Females manipulated spermatophores received from typus males for long time periods (minutes), but not those they received from robustus males. By accepting sperm from subordinate typus males, females may avoid further harassment (convenience polyandry), but they subsequently may discriminate against these subordinate males by delaying spawning and removing their sperm. These observations suggest that female behavior influences the outcome of matings, favoring fertilization of eggs by sperm from dominant males. Convenience polyandry and cryptic female choice may be common in other crustaceans as well.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by T. Czeschlik     

Food site residence time and female competitive relationships in wild gray-cheeked mangabeys (<Emphasis Type="Italic">Lophocebus albigena</Emphasis>)

Authors of socioecological models propose that food distribution affects female social relationships in that clumped food resources, such as fruit, result in strong dominance hierarchies and favor coalition formation with female relatives. A number of Old World monkey species have been used to test predictions of the socioecological models. However, arboreal forest-living Old World monkeys have been understudied in this regard, and it is legitimate to ask whether predominantly arboreal primates living in tropical forests exhibit similar or different patterns of behavior. Therefore, the goal of our study was to investigate female dominance relationships in relation to food in gray-cheeked mangabeys (Lophocebus albigena). Since gray-cheeked mangabeys are largely frugivorous, we predicted that females would have linear dominance hierarchies and form coalitions. In addition, recent studies suggest that long food site residence time is another important factor in eliciting competitive interactions. Therefore, we also predicted that when foods had long site residence times, higher-ranking females would be able to spend longer at the resource than lower-ranking females. Analyses showed that coalitions were rare relative to some other Old World primate species, but females had linear dominance hierarchies. We found that, contrary to expectation, fruit was not associated with more agonism and did not involve long site residence times. However, bark, a food with a long site residence time and potentially high resource value, was associated with more agonism, and higher-ranking females were able to spend more time feeding on it than lower-ranking females. These results suggest that higher-ranking females may benefit from higher food and energy intake rates when food site residence times are long. These findings also add to accumulating evidence that food site residence time is a behavioral contributor to female dominance hierarchies in group-living species.     

Sex differences in fitness following a group take-over among Toque macaques: testing models of social evolution

Summary A group of toque macaques took-over the home range of one of its subordinate neighboring groups and fused with it to form a larger cohesive group. In the 7 years before the take-over, the dominant group had consistently won all contests at common feeding sites, yet the fitnesses of the females of these two groups did not differ significantly (Fig. 2A). After the take-over the females of the subjugated group occupied the lowest ranks in the combined dominance hierarchy of the merged groups (Fig. 1) and thereby lost the advantages of an own home range, such as priority of access to food. Consequently, in the merged group, survivorship and reproductive success among the subjugated females were significantly less than among the females of the dominant subgroup (Table 2, 4). The dominant matrilines grew numerically and replaced all of the subjugated females, and all but one of their offspring, within 8 years after the take-over (Fig. 2B). These data support the hypothesis that cooperation among female kin in defending resources against strange females is important in the evolution of female-bonded groups. Before the merger all 5 natal males of the subordinate group had transferred to the dominant group, where they occupied high and mid-level dominance ranks (Fig. 1). These males survived at a significantly greater rate than their subordinate female kin. Thus, the cost of group transfer seems to be greater for females than for males, and this may be one reason that females generally do not emigrate or that groups do not fuse. The data suggested three hypotheses. First, since large body size and other adaptations for fighting, giving males an advantage in male-male competition for mates, are also of advantage in resource competition with males and females, such male characters may also be favored by non-sexual selection, especially where male reproductive strategy involves group transfer. Second, female bonded groups evolved as female defensive coalitions against not only female but also male resource competitors, there having been a mutual influence in the coevolution of large-sized males and female gregariousness. Third, female defensive coalitions against large-sized aggressive males are also advantageous out-side the context of food competition, or, independent of foraging strategy.     

Physiological traits associated with a reproductive-resting stage in Coullana canadensis (Copepoda: Harpacticoida)

We investigated physiological traits associated with a reproductive-resting stage in adult female harpacticoid copepods, Coullana canadensis (Willey). Our hypothesis was this stage represents a life-history strategy that could increase fitness by improving winter survivorship and future reproductive success in spring. To test if physiological rates are suppressed in this stage, we compared gut-cell morphology, nitrogen excretion rates, enzyme activities, and phytoplankton grazing rates of reproductive-resting and reproductive females reared in the laboratory under high food conditions. Copepods came from laboratory cultures originating from individuals collected in Maine and Maryland, USA in 1990. Reproductive-resting females had lower physiological rates, and the surface area of gut-cells was reduced compared to reproductive females. Distinct morphological differences in the distribution of lipids between reproductive and reproductive-resting females were observed under light and electron microscopy, the latter having a diffuse accumulation of lipid in the area normally occupied by the ovaries. Differences in lipid composition were also found. Reproductive copepods had a significantly higher ratio (%) of polar lipids to total lipid, and a lower proportion of triacylglycerols compared to reproductive-resting copepods. These laboratory findings are consistent with the hypothesis that the reproductive-resting stage in C. canadensis is an adaptive response to increase winter survival. Received: 29 May 1996 / Accepted: 3 December 1997     

No evidence of mate discrimination against males carrying a sex ratio distorter in <Emphasis Type="Italic">Drosophila pseudoobscura</Emphasis>

Selfish genetic elements (SGEs) that spread by manipulating spermatogenesis often have highly deleterious effects on males that carry them. Females that mate with male carriers of SGEs can also suffer significant costs: they receive fewer and poorer-quality sperm, their offspring will inherit the deleterious allele, and the sex ratio of their offspring will be biased towards the more common sex. To counter these costs, females are therefore expected to prefer to mate with males that do not carry sex ratio distorters or other deleterious selfish genetic elements. However, despite the potential costs, there are few examples of female choice against males carrying SGEs. We searched for evidence of a female preference in fruit fly Drosophila pseudoobscura against males carrying a costly meiotic driving X-chromosome Sex Ratio (SR). In a series of five non-competitive mate preference experiments, we find no evidence that females prefer to mate with non-SR males. Our use of five separate experiments, involving more than 800 females, makes it unlikely that this lack of a difference was due to low power or simple chance. We suggest that the lack of female choice against SGE-carrying males may be due to strong selection on SGEs to be indistinguishable from alternative alleles. Furthermore, polyandry, either in direct response to receiving an ejaculate from an SGE-carrying male or carried out indiscriminately when at risk of mating with carriers, may be an alternative response by females to limit the exposure of their offspring to SGEs.     

Reproductive trade-offs from mating with a successful male: the case of the tephritid fly Anastrepha obliqua

In lekking species, females may become sperm-limited when mating with sexually successful males, and this may be exacerbated by a poor male diet. Polygynous males may also be limited by the amount of accessory gland products (AGPs) they can transmit to females, which in turn may influence the females’ refractory period and longevity. Here, we tested the effect of male mating history, larval and adult diet on copula duration, mating intervals, female fecundity, fertilisation success, life span and likelihood to remate using sexually successful males of the lekking tephritid fly Anastrepha obliqua. Flies originated from either a native or exotic host fruit and were protein-fed or deprived. Male diet and larval host influenced copula duration, while the time elapsed between matings was affected by the interaction of mating order and male adult diet. Female fecundity was not influenced by female position in mating order or protein inclusion into the male diet. However, mating order and male larval diet influenced female fertilisation success. Importantly, as males mated successively they were less able to induce a refractory period on females, as the last females to mate with a male were more likely to remate and had slightly longer life spans than the first females to mate with males. These results might be attributed to a decrease in male AGPs with increasing male mating frequency. We discuss the role of conditional expression of male mating frequency with respect to A. obliqua’s life history, the trade-off that females face when mating with a successful male, the effect of larval diet on adult sexual performance and the possibility for sexual conflict to occur due to high male mating rates and fitness costs to females.     

The evolution of empty nuptial gifts in a dance fly, Empis snoddyi (Diptera: Empididae): bigger isn't always better

Adaptive female choice is thought to have led to the evolution of nutritionally valuable nuptial gifts in many insect species. However, in several dance fly species, males offer and females accept “empty gifts” with no nutritional value. In the species studied here, Empis snoddyi Steyskal, males produce empty balloons comprised of hundreds of silk bubbles and form mating swarms that females approach to investigate males. Males within the swarm engage in agonistic interactions. The empty balloon has been hypothesized to be an indicator of male condition such that males with larger balloons are predicted to have higher mating success and be more successful in male-male interactions than males with smaller balloons. We examined the role of male body size and balloon size in the context of intersexual and intrasexual selection. We found that neither male body size nor balloon size affected the outcome of pairwise male-male interactions. Using multiple-regression techniques, we found significant linear selection for increasing male body size and decreasing balloon size associated with mating success, a surprising result given a positive relationship between male body size and balloon size. A visualization of selection showed the highest peak of male mating success for larger males with intermediate-size balloons. These results can be explained by a trade-off between long-range attraction of females using large balloons and close-range attraction of females via improved flying efficiency associated with smaller balloons. Both male body size and balloon size are important components in determining male mating success; however, the empty balloon does not appear to play a typical role as a sexually selected ornament. Received: 29 December 1997 / Accepted after revision: 7 October 1998     

Copulation duration and sperm precedence in the stalk-eyed fly Cyrtodiopsis whitei (Diptera : Diopsidae)

Summary By means of field observations and laboratory experiments on the Malaysian stalk-eyed fly Cyrtodiopsis whitei we examined the consequences of variation in copulation duration for sperm competition. In this sexually dimorphic species over 90% of all copulations occur in nocturnal aggregations with from one to four males and up to 24 females. Copulation duration observed in both the field and the laboratory exhibited a bimodal distribution with peaks at 10 and 50 s. In the field short copulations less than 30 s long occurred frequently when more than one male was present in an aggregation but most were not the direct result of male interference. Sperm counts from female spermathecae after artificial interruptions indicated sperm are not transferred during the first 40 s of a copulation. When solitary males mated up to five times in succession to virgin females, short copulations did not occur, nor was the number of sperm transferred reduced. However, short copulations did occur when we mated isolated females within 6 min of a previous copulation. By mating irradiated and non-irradiated males in reciprocal pairs we discovered that C. whitei exhibits both first-male sperm precedence and sperm mixing. More than half of the females mated first to sterile and then to fertile males failed to produce offspring. Such variation in copulation duration and sperm precedence is consistent with male placement and detection of a spermatophore that acts as a temporary mating plug. Our data suggest that those male C. whitei which successfully defend large aggregations of females reduce sperm waste and competition by preferentially transferring sperm to females that have not mated recently. Correspondence to: G.S. Wilkinson     

Effect of female molt stage and sex ratio on courtship behavior of the blue crab Callinectes sapidus

In many species, males and females actively participate in courtship, and the outcome of pre-mating interactions influences the mating success of both sexes. Female blue crabs, Callinectes sapidus, mate soon after their final molt to maturity; thus female molt stage dictates the timing of mating. In a field experiment, we manipulated female molt stage and sex ratio to test their effects on the courtship behavior of both sexes, if female behavior influences the behavior and pairing success of males, and if male courtship influences male pairing-success. Early-molt-stage females avoided males during courtship, whereas late-molt-stage females sought out males. As a result, males had to pursue and capture early-molt-stage females whereas males displayed to late-molt-stage females and more easily physically controlled them. Males sometimes abandoned late-molt-stage females, but this occurred more often when females were abundant. The rate at which females avoided males was positively correlated with that of males abandoning females, and males that were unsuccessful at pairing met with higher rates of female resistance than successful males, suggesting that female behavior influences male pairing-success. Unlike unsuccessful males, successful males more often made the transition between display and maintaining physical control of the female. At high male sex ratios, males initiated courtship more readily; thus both sexual competition and female behavior influence male courtship in this species. Received: 7 July 1996 / Accepted: 10 January 1998     

Non-independent mate choice in a fiddler crab: a case of stimulus enhancement

Non-independent mate selection occurs when the choice behavior of a female is altered by the interactions between other females and males. In the fiddler crab Uca mjoebergi, males court mate-searching females by waving their one greatly enlarged claw. When a female approaches a male, he initiates high-intensity waving. We conducted one natural mate choice experiment and two mate choice experiments using custom-built robotic crabs. We show that the decision of one female to approach a group of males increases the probability that another female will approach and visit a male from the same group. We suggest that this behavior is best explained by the ‘stimulus enhancement’ hypothesis, where the presence of a female near a group of males makes them more likely to be detected by other females due to an increase in male display rate.     

Testing function of fiddler crab claw waving by manipulating social context

Many territorial advertisement signals are thought to be dual-function signals, directed to both rival male and receptive female conspecifics. However, few studies have tested this assumption by examining whether in fact both sexes are likely to elicit signaling behavior from territorial males. In this study, I experimentally manipulated the social context of male sand fiddler crabs (Uca pugilator) to investigate the effect of different audiences on the performance of the claw-waving display, a territorial signal that is often presumed to be directed to both males and females. To test whether males perform this signal to both audiences, I measured the frequency of waving behavior by focal males when housed in field enclosures alone, with only males, with only females, or with both males and females. Focal males waved at a low frequency when alone, and the presence of males had no effect on their level of waving. However, in the presence of females, focal males showed a significantly higher level of waving, whether or not males were also present. In addition, there was no association between fighting and waving behavior. This experiment provides evidence that from the perspective of the signaling male, the claw-waving display of U. pugilator is not a dual-function signal but rather is primarily directed to receptive females. Received: 16 December 1999 / Received in revised form: 1 February 2000 / Accepted: 19 February 2000     

Determinants of male spacing behavior in microtines: an experimental manipulation of female spatial distribution and density

Theory on microtine mating systems predicts that male spacing behavior will be related to female spatial and temporal distribution. However, data from a natural population of field voles, Microtus agrestis, indicated a potential influence of female density on the spacing behavior of males. Therefore, I experimentally investigated the relative importance of female density and female spatial distribution for the spacing behavior of males in M. agrestis. Males were radio-tracked in enclosed natural habitats in which females at different densities were placed in two different spatial arrangements: clumped versus even distribution. Female density was the main factor determining male spacing behavior. At the high female density males had smaller home ranges and moved shorter distances between radio-tracking recordings. Also, home ranges were more exclusive at high female density. However, since there was a significant positive correlation between home range size and range overlap, range exclusiveness seemed to be influenced by female density indirectly through the effect of range size. Female spatial distribution, on the other hand, had no influence on male home range exclusiveness.     

Inter-male competition for high-quality host-plants: the evolution of protandry in a territorial grasshopper

Summary Males of the desert grasshopper Ligurotettix coquilletti typically eclose 3–5 weeks prior to females. Early-eclosed males experience more female encounters than the later-eclosed individuals. Evidence suggests that the number of encounters may be proportional to male lifetime mating success. Early-eclosed males enjoyed greater adult lifespans, occupied and defended higher quality territories (Larrea shrubs), and tended to be dominant on these shrubs if several males were present. The elevated number of female encounters were not conferred on early-eclosed males by their extended survivorship, but rather by the high quality of their territories, which retained numerous females, and their dominant behavior. We evaluate several female-benefit and male-benefit hypotheses for the evolution of protandry and propose that in L. coquilletti, the phenomenon arose due to male-male competition for female-encounter sites, a mechanism not considered in earlier models. Finally, we discuss several reasons for the high variance in the date of male eclosion.     

Parasitoid learning during interactions with ants: how to deal with an aggressive antagonist

Pauesia picta, P. pinicollis and P. silvestris (Hymenoptera: Aphidiidae) are common parasitoids of the conifer aphid Cinara pinea, which is regularly attended by red wood ants (Formica spp.). In this study, I tested whether females of these parasitoid species learned during interactions with honeydew-collecting Formica polyctena workers that caution is not necessary if searching behaviour is adapted, and whether parasitoids benefit from being able to learn. When searching on Scots pines, naive females of P. picta and P. pinicollis generally retreated to a pine needle when making contact with a honeydew-collecting ant, did not approach ants from the side or from the front and kept a ”safe distance” from ants when sitting on needles. After some non- aggressive ant encounters, experienced female parasitoids changed their behaviour: they reduced their searching speed, approached ants from the side and even from the front, retreated less often in response to an approaching ant and reduced the ”safe distance”. These experienced females had a significantly higher rate of oviposition than naive females or females foraging for an unattended host. Thus, the ability of the parasitoid to learn during interactions with an antagonist led to a prolonged retention time and a higher oviposition rate. By contrast, there was no evidence of learning in P. silvestris. Females of this species showed no behavioural change in response to ant encounters, and there was no difference in the foraging success of naive and experienced female parasitoids. Received: 7 December 1999 / Revised: 23 September 2000 / Accepted: 10 March 2000     

Female encounter rates and fighting costs of males are associated with lek size in Drosophila mycetophaga

Male costs and benefits associated with male display size in field populations of an Australian lekking Drosophila species were examined. Results suggested that male mating success increased with display size, since matings appeared to be more common in large displays, and since the probability of males encountering a female increased as displays contained more males. Female encounter probabilities did not increase once about 20 males or more were present on a display. Male size and fighting costs tended to increase with display size. The distribution of males among displays did not follow the ideal free distribution in the sense that each male did not have equal mating opportunity per unit time. Deviation from an ideal free distribution may have been due to female preference for mating in aggregations rather than with solitary males, since in a field experiment females were more willing for mating in an aggregation of five males than with solitary males. Received: 22 May 1997 / Accepted after revision: 1 November 1997